A new species of the genus Arachnothelphusa Ng, 1991 (Crustacea: Decapoda: Gecarcinucidae) from a limestone cave in Sarawak (Malaysian Borneo) ...
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RAFFLES BULLETIN OF ZOOLOGY 2021
Taxonomy & Systematics
RAFFLES BULLETIN OF ZOOLOGY 69: 1–7
Date of publication: 12 January 2021
DOI: 10.26107/RBZ-2021-0001
http://zoobank.org/urn:lsid:zoobank.org:pub:F861B9E5-475A-4A3D-B2AC-B49646FA7CC9
A new species of the genus Arachnothelphusa Ng, 1991 (Crustacea:
Decapoda: Gecarcinucidae) from a limestone cave in Sarawak
(Malaysian Borneo)
Jongkar Grinang1* & Peter K. L. Ng2
Abstract. A new species of cavernicolous gecarcinucid crab, Arachnothelphusa sarang, is described from a
limestone cave in northern Sarawak, Malaysian Borneo. This increases the number of Arachnothelphusa species
to six. It is the second member in the genus that is known to primarily occupy limestone caves, the other being
A. rhadamanthysi Ng & Goh, 1987, from Gomantong in Sabah. Both species appear to be cavernicolous species
with pale body colouration in life.
Key words. Brachyura, taxonomy, Oriental region, freshwater crab, cavernicolous crab
INTRODUCTION Chian National History Museum (former Raffles Museum
of Biodiversity Research), National University of Singapore
Currently, the Bornean gecarcinucid genus Arachnothelphusa (ZRC); Sarawak Biodiversity Centre, Sarawak, Malaysia
Ng, 1991, is represented by five species, viz. A. melanippe (SBC); Naturalis Biodiversity Center (former Rijksmuseum
(De Man, 1899) [central Kalimantan], A. kadamaiana van Natuurlijke Historie), Leiden, The Netherlands (RMNH);
(Borradaile, 1900) [northern Sabah], A. rhadamanthysi and Senckenberg Museum und Forschungsinstitut, Frankfurt
Ng & Goh, 1987 [eastern Sabah], A. terrapes Ng, 1991 am Main, Germany (SMF).
[eastern Sabah], and A. merarapensis Grinang, Pui & Ng,
2015 [northern Sarawak] (Grinang et al., 2015; Ng & Ng,
2018). Members of Arachnothelphusa live in a wide range TAXONOMY
of habitats, from tree-holes to the interior of limestone
caves. Of the five species, only A. rhadamanthysi has been Family Gecarcinucidae Rathbun, 1904
recorded from limestone caves in Gomantong in Sabah.
We here describe a sixth species of Arachnothelphusa, A. Arachnothelphusa Ng, 1991
sarang, new species, and the second cavernicolous member
from a limestone cave system in Sarawak. Type species. Potamon (Potamon) melanippe De Man, 1899,
by original designation.
MATERIAL AND METHODS Arachnothelphusa sarang, new species
(Figs. 1A–F, 2A–G, 3A–E, 4A)
The terminology used essentially follows Ng (1988) and
Davie et al. (2015), with the abbreviations G1 and G2 Material examined. Holotype: male (20.4 × 14.7 mm)
used for the male first and second gonopods, respectively. (ZRC 2020.0098), limestone cave, Bukit Sarang, Bintulu,
Measurements provided in millimetres are of the carapace Sarawak, Malaysia, coll. H.H. Tan et al., 20 August 2005.
width and length, respectively. Specimens examined are Paratypes: 1 male (18.7 × 14.8 mm), 4 females (15.8–19.8
deposited in the Zoological Reference Collection, Lee Kong × 12.0–15.8 mm) (ZRC 2020.0099), same data as holotype;
10 males (7.4–11.2 × 5.8–9.6 mm), 7 females (7.5–12.7 ×
5.8–9.9 mm) (ZRC 2020.0100), limestone cave, Batu Gelam,
Bukit Sarang, Bintulu, Sarawak, Malaysia, coll. H.H. Tan,
Accepted by: Jose Christopher E. Mendoza 20 August 2005; 1 male (12.1 × 9.9 mm), 1 female (12.9 ×
10.4 mm) (ZRC 2020.0351), limestone cave, Batu Kelelut,
1
Institute of Biodiversity and Environmental Conservation, Universiti Malaysia
Sarawak, 94300 Kota Samarahan, Sarawak, Malaysia; Email: gjongkar@unimas.my Bukit Sarang, Bintulu, Sarawak, Malaysia, coll. H.H. Tan
(*corresponding author) et al., 18 August 2005.
2
Lee Kong Chian Natural History Museum, Faculty of Science, National University
of Singapore, 2 Conservatory Drive, Singapore 117377, Republic of Singapore
Comparative material. Arachnothelphusa merarapensis
© National University of Singapore Grinang, Pui & Ng, 2015: Holotype male (22.5 × 16.8
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print) mm) (ZRC 2016.0297), water-filled tree-hole, ca. 100
1
Grinang & Ng: New species of Arachnothelphusa from limestone cave in Sarawak
Fig. 1. Arachnothelphusa sarang, new species. A, male (12.1 × 9.9 mm) (ZRC 2020.0351), Batu Rusa cave, Bukit Sarang, Bintulu,
Sarawak, Malaysia; B, female (12.9 × 10.4 mm) (ZRC 2020.0351), Batu Kelelut, Bukit Sarang, Bintulu, Sarawak, Malaysia, specimen;
C–E, paratype male (18.7 × 15.3 mm) (ZRC 2020.0099), Bukit Sarang, Bintulu, Sarawak, Malaysia; F, paratype female (15.8 × 11.8 mm)
(ZRC 2020.0099), Bukit Sarang, Bintulu, Sarawak, Malaysia. A, B, photographed in situ; C, F, overall dorsal habitus; D, ventral view of
cephalothorax; E, frontal view of cephalothorax and chelae. Photographs: Tan Heok Hui.
cm above ground, steep dipterocarp forest, Merarap Hot × 17.1 mm) (ZRC 2009.0094), Poring, Basin 1A, Sabah,
Spring Resort, Lawas, northern Sarawak, Malaysia, Borneo, Borneo, coll. R.F. Inger et al., 12 August 1992; 3 males
4°22′25.4″N 115°26′10.1″E, 485 m asl, coll. J. Grinang & (21.1 × 15.8 mm, 22.8 × 16.5 mm, 25.3 × 18.5 mm) (ZRC
Y.M. Pui, 31 October 2014; paratype female (19.9 × 15.2 2002.0097), Crocker Range, Sabah, 5°27′N 116°03′E, coll.
mm) (SBC.C.00376), water-filled hole of tree buttress, ca. I. Das, 24 April 2001. Arachnothelphusa aff. kadamaiana: 1
90 cm above ground, steep dipterocarp forest, Merarap Hot female (19.0 × 14.2 mm) (ZRC 2002.0098), Bako National
Spring Resort, Lawas, northern Sarawak, Malaysia, Borneo, Park, Sarawak, coll. I. Das & L. Grismer, 27 March 2001.
4°22′16.5″N 115°26′12.4″E, 494 m asl, coll. J. Grinang & Arachnothelphusa terrapes Ng, 1991: Holotype male (17.6
Y.M. Pui, 1 November 2014; 1 female (22.3 × 16.1 mm) × 13.3 mm) (ZRC 1992.7918), Danum Valley Field Centre,
(SBC.C.00377), water-filled hole of tree buttress, ca. 30 cm station 507, in dry stump on ridge, Lahad Datu, Sabah,
above ground, same data as paratype, coll. Y.M. Pui, 27 Borneo, leg. H.K. Voris, 23 October 1990; paratype female
February 2013. Arachnothelphusa kadamaiana (Borradaile, (25.7 × 18.6 mm) (ZRC 1992.7919), Danum Valley, Lahad
1900): Holotype female (18.9 × 13.5 mm) (SMF 4281), Datu, Sabah, Borneo, leg. S.C. Choy, 21 July 1989; others:
Kadamian River, Sabah, Malaysia, Borneo; 1 male (20.1 × 1 male (30.8 × 20.5 mm), 1 female (30.1 × 20.5 mm, with
14.9 mm) (SMF 4282), same data as holotype; 1 female (23.2 26 juvenile crabs) (ZRC 2017.1205), from water-filled tree
2RAFFLES BULLETIN OF ZOOLOGY 2021
Fig. 2. Arachnothelphusa sarang, new species. A–E, holotype male (20.4 × 14.7 mm) (ZRC 2020.0098), Bukit Sarang, Bintulu, Sarawak,
Malaysia; F, G, paratype female (19.8 × 15.8 mm) (ZRC 2020.0100), Bukit Sarang, Bintulu, Sarawak, Malaysia. A, F, dorsal view of
habitus; B, G, ventral view of carapace showing pleon; C, frontal view of cephalothorax; D, outer view of left chela; E, outer view of
right fourth ambulatory leg. Scales: A, B, C, F, G = 10 mm; D, E = 5 mm.
buttress, ca. 35 cm above ground Danum Valley, Lahad Datu, Females. The females differ in minor non-sexual characters
Sabah, Borneo, Malaysia, 20 July 2017. Arachnothelphusa by the carapace being slightly broader and lower (Fig. 2F,
melanippe (De Man, 1899): Lectotype male (18.9 × 14.4 mm) G). Female pleon broad, round, somite 6 subequal to length
(RMNH D1303a), Liang Koebeng Mountains, Kalimantan, of telson, tip of telson round pointed (Fig. 2G). In larger
leg. 1897; paralectotype female (21.4 × 16.7 mm) (RMNH specimens of both sexes, the antero- and posterolateral
D1303b), same as lectotype. regions have relatively more coarse granules. The vulvae
are transversely ovate, large, without obvious sternal vulvar
Diagnosis. Carapace surface convex, rugose, finely covers and are positioned on the median part thoracic of
granular; anterolateral margins convex, serrated; antero- and sternite 6.
posterolateral regions prominently rugose, covered with
numerous coarse granules; epibranchial tooth very low or Variation. The numerous paratype specimens agree well
indistinct; external orbital tooth very low, broadly triangular, with the holotype male in non-sexual characters.
outer margin slightly concave, distinctly serrated; epigastric
and postorbital cristae distinct; cervical and H-grooves deep, Colour. In life, the species is pale purplish brown to yellowish
not confluent (Fig. 2A, C); ambulatory legs long, merus of in both sexes on the dorsal surfaces, the ventral surfaces
fourth ambulatory legs subequal to length of carapace (Fig. being pale yellow to dirty white. The corneas of the eyes
2A, B, E); carpus of chelipeds rugose, with fine granules, are large and fully pigmented (Fig. 1).
inner angle with broadly triangular tooth (Fig. 2A, D); chela
relatively short, fingers as long as palm, cutting teeth on Etymology. The species is named after the locality where
pollex not prominent (Fig. 2D). Male pleon T-shaped, somite the holotype was collected. The name is used as a noun in
6 subequal to length of telson (Fig. 2B). G1 slender, sinuous, apposition.
gently curving outwards; terminal segment cylindrical,
tapering, about one third length of subterminal segment (Fig. Remarks. Arachnothelphusa sarang, new species, is
3A–D). G2 with short distal segment, less than a quarter easily distinguished from A. merarapensis, A. terrapes,
length of basal segment (Fig. 3E). A. melanippe, and A. rhadamanthysi by its very low to
indistinct epibranchial tooth as well as the very low, broad
3Grinang & Ng: New species of Arachnothelphusa from limestone cave in Sarawak
Fig. 3. Arachnothelphusa sarang, new species, holotype male (20.4 × 14.7 mm) (ZRC 2020.0098), limestone cave, Bukit Sarang, Bintulu,
Sarawak, Malaysia. A, dorsal view of left G1; B, dorsal view of distal part of left G1; C, ventral view of left G1; D, ventral view of distal
part of left G1; E, dorsal view of left G2. Scales = 0.5 mm.
external orbital tooth (Figs. 1C, F, 2A–C, F, G, 4A). In A. 3A–D, 5K–M); while that of A. merarapensis is about half
merarapensis, the epibranchial tooth is acutely triangular, the length of the subterminal segment (Fig. 5A–D). The
separated from the external orbital tooth by a wide and deep distal segment of the G2 of A. sarang and A. merarapensis
cleft and the external orbital tooth is acutely triangular (Fig. is less than a quarter the length of the basal segment (Figs.
4B; cf. Grinang et al., 2015: fig. 1A–C). In A. terrapes, the 3E, 5E), but in A. kadamaiana, it is less than one-fifth the
epibranchial tooth is distinct on both sides, separated from length of the basal segment (Fig. 4N). The gonopods of A.
the external orbital tooth by a deep and broad U-shaped cleft rhadamanthysi are not known.
and the external orbital tooth is triangular (Fig. 4C; cf. Ng,
1991: fig. 3 [incorrectly printed as fig. 5]; cf. Ng & Ng, 2018: The live colouration of the two cave species differs markedly
fig. 5B–F). In A. melanippe, the epibranchial tooth is distinct from those of the epigeal species. The live colouration of A.
but relatively small and separated from the external orbital sarang, new species, ranges from light purplish to yellowish
tooth by a small cleft, with the external orbital tooth broadly brown (Fig. 1). This is similar to that known for the other
triangular (Fig. 4E; cf. De Man, 1899: pl. 9 fig. 11; Ng, cavernicolous species, A. rhadamanthysi, which is pale
1991: fig. 1). In A. rhadamanthysi, the epibranchial tooth is straw-yellow with white legs (Fig. 6; cf. Ng & Goh, 1987:
distinct on both sides with the external orbital tooth triangular 326). Arachnothelphusa merarapensis is bright purple overall
(cf. Ng & Goh, 1987: pl. 3A, B). In carapace features, A. (cf. Grinang et al., 2015: fig. 3A, B), whereas A. terrapes is
sarang is perhaps closest to A. kadamaiana, but they can brown to reddish-brown (cf. Ng, 1991: figs. 4, 5 [not labelled
be distinguished by the presence of a distinct epibranchial in original paper]; Ng & Ng, 2018: fig. 5B–E). The habits
tooth and a relatively more anteriorly projecting and acute and live colouration of A. melanippe and A. kadamaiana
apex of the external orbital tooth for the latter (Figs. 2A–C, are not known.
F, G, 4A versus Fig. 4D; cf. Grinang et al., 2015: fig. 6A).
The anterolateral region of A. sarang is distinctly less rugose The habitat, pale colouration in life, and general appearance
with fine granules, whereas this region is prominently rugose of Arachnothelphusa sarang, new species, are also similar
with coarse granules for congeners (Fig. 4A versus 4B–E). to those of the genus Stygothelphusa Ng, 1989, which
The structure of the G1 of A. sarang differs from that of A. contains four species, all from limestone caves in Sarawak
terrapes and A. melanippe in gently curving outwards (Fig. (Ng, 1991, 2013; Ng & Grinang, 2014). The carapace
3A–D) versus distinctly curving outwards in the latter two of Stygothelphusa, however, is generally more quadrate
species (Fig. 5F–I, O–R). In the G1 structure, A. sarang is (versus more transversely ovate in Arachnothelphusa), the
similar to A. merarapensis and A. kadamaiana, but they ambulatory legs are proportionately even longer, and most
can be distinguished by the following features: the terminal significantly, the G2 is much longer than the G1, with the
segment of the G1 of A. sarang and A. kadamaiana is distal segment long and flagelliform (versus G2 shorter than
about one-third the length of the subterminal segment (Figs. G1, with the distal segment short in Arachnothelphusa).
4RAFFLES BULLETIN OF ZOOLOGY 2021
Fig. 4. Carapace morphology of Arachnothelphusa species. A, A. sarang, new species, holotype male (20.4 × 14.7 mm) (ZRC 2020.0098),
limestone cave, Bukit Sarang, Bintulu, Sarawak; B, A. merarapensis Grinang, Pui & Ng, 2015, holotype male (22.5 × 16.8 mm) (ZRC
2016.0297), Merarap Hot Spring, Lawas, Sarawak; C, A. terrapes Ng, 1991, male (30.8 × 20.5 mm) (ZRC 2017.1205), Danum Valley,
Lahad Datu, Sabah; D, A. kadamaiana (Borradaile, 1900), male (20.1 × 14.9 mm) (SMF 4282), Kadamian River, Sabah; E, A. melanippe
(De Man, 1899), paratype female (21.4 × 16.7 mm) (RMNH D1303b), Mt. Liang Koebeng, Kalimantan, Indonesia. E after Ng (1991).
Fig. 5. Gonopod morphology of Arachnothelphusa species. A–E, A. merarapensis Grinang, Pui & Ng, 2015, holotype, male (22.5 ×
16.8 mm) (ZRC 2016.0297), Merarap Hot Spring, Lawas, Sarawak; F–J, A. terrapes Ng, 1991, holotype, male (17.6 × 13.3 mm) (ZRC
1992.7918), Danum Valley, Lahad Datu, Sabah; K–N, A. kadamaiana (Borradaile, 1900), male (20.1 × 14.9 mm) (SMF 4282), Kadamian
River, Sabah; O–S, A. melanippe (De Man, 1899), lectotype, male (18.9 × 14.4 mm) (RMNH D1303a), Mt. Liang Koebeng, Kalimantan,
Borneo. C, F, K, O, ventral view of left G1; D, G, L, P, ventral view of distal part of left G1; A, H, Q, dorsal view of left G1; B, I, M,
R, ventral view of distal part of left G1; E, J, N, S, left G2. A–E, K–N after Grinang et al. (2015); F–J, O–S after Ng (1991).
5Grinang & Ng: New species of Arachnothelphusa from limestone cave in Sarawak
hill surrounded by peat swamp, comprises a complex of
small caves with different names, most of which are almost
certainly with subterranean interconnections, and is part of
the Tatau River basin. All the specimens of A. sarang were
found in water pools with fine substrates, and co-existing
with whelk snails, several hundred metres from the cave
entrance; none were observed near the cave mouth (H. H.
Tan, pers. comm.). The limestone cave of Bukit Sarang is
a protected area and important for sustainable edible bird-
nest production. The harvesting of bird-nests will have to
be done carefully so as not to pollute or disturb the habitat
if the crab is to be conserved.
Key to species of Arachnothelphusa of Borneo
1. Epibranchial tooth very low or indistinct; external orbital tooth
very low, broad; anterolateral region less rugose, fine granules,
carapace surface smooth (Figs. 1C, F, 2A, B, C, F, G, 4A);
G1 gently curving outwards, terminal segment about one-third
length of subterminal segment; distal segment of G2 less than
a quarter length of basal segment (Fig. 3A–E).........................
............................ A. sarang, new species (northern Sarawak)
– Epibranchial tooth distinct, triangular; external orbital tooth
triangular; anterolateral region prominently rugose, coarse
granules, carapace surface rugose (Fig. 4B–E); G1 gently or
strongly curving outwards, terminal segment either half, one-
third or a quarter length of subterminal segment; distal segment
Fig. 6. Live colours of Arachnothelphusa rhadamanthysi from of G2 either a quarter or one-fifth length of basal segment (Fig.
Gomantong limestone cave in Sabah, specimen not collected. A, 5)...............................................................................................2
male, from outside of the cave; B, inside of cave. Photographs: 2. Epibranchial tooth distinct, acutely triangular; G1 strongly
Keith Christenson. curving outwards (Fig. 5F, H, O, Q).......................................3
– Epibranchial tooth distinct, relatively blunt; G1 gently curving
outwards (Fig. 5A, C, K).........................................................4
Members of Arachnothelphusa occur in a wide variety
3. Epibranchial tooth acutely triangular, distinctly separated from
of habitats ranging from tree-holes to caves. The only external orbital tooth by wide, deep cleft; external orbital
other species of Arachnothelphusa known from caves acutely triangular (Fig. 4B); terminal segment of G1 about
is A. rhadamanthysi. Arachnothelphusa sarang inhabits half of length of subterminal segment; distal segment of G2
exclusively deep interior of cave systems of Bukit Sarang about quarter length of basal segment (Fig. 5A–E)..................
in northern Sarawak. Arachnothelphusa rhadamanthysi on ....................................... A. merarapensis (northern Sarawak)
the other hand, has been found in more exposed areas near – Epibranchial tooth distinct but relatively small, separated from
the cave entrance of Gomantong limestone cave in western external orbital tooth by small cleft; external orbital tooth
Sabah (Fig. 6). Both species, however, are not troglobitic broadly triangular (Fig. 4E); terminal segment of G1 about
one-third of length of subterminal segment; distal segment
species as neither have their eyes reduced and the corneas
less than a quarter length of basal segment (Fig. 5O–S)..........
are still completely pigmented, and A. rhadamanthysi at least .......................................... A. melanippe (central Kalimantan)
has been found at cave entrances. In this respect, the habits 4. Epibranchial tooth distinct; external orbital tooth broadly
of the two species are the same as species of Stygothelphusa, triangular (Fig. 4D); terminal segment of G1 about a third
which are regarded as troglophilic taxa instead (Ng, 1991, length of subterminal segment (Fig. 5K–M).............................
2013; Ng & Grinang, 2014). Another cavernicolous ............................................. A. kadamaiana (northern Sabah)
gecarcinucid from Borneo, Balssiathelphusa phasma Ng & – Epibranchial tooth distinct; external orbital tooth triangular;
Guinot, 2014, from eastern Kalimantan in Indonesia, is also terminal segment of G1 about a quarter length of subterminal
regarded as troglophilic (Ng & Guinot, 2014). As such, the segment.....................................................................................5
5. Epibranchial tooth distinct on both sides, separated from
only true stygobitic crabs in Borneo are the two species of
external orbital tooth by deep, broad U-shaped cleft; external
Cerberusa Holthuis, 1979, from northern Sarawak (family orbital tooth triangular (Fig. 4C); G1 with cone-shaped terminal
Potamidae) (Holthuis, 1979) and the monotypic Guaplax segment, about a quarter length of subterminal; distal segment
Naruse, Ng & Guinot, 2008, from southern Kalimantan of G2 less than fifth length of basal segment (Fig. 5F–J)........
(family Hymenosomatidae) (Naruse et al., 2008). ...................................................... A. terrapes (eastern Sabah)
– Epibranchial tooth distinct on both sides, small, blunt, not
Habitat. The habitat of A. sarang, new species, is a limestone separated from external orbital tooth by cleft...........................
outcrop. Bukit Sarang in Bintulu is an isolated limestone ........................................... A. rhadamanthysi (eastern Sabah)
6RAFFLES BULLETIN OF ZOOLOGY 2021
ACKNOWLEDGEMENTS Naruse T, Ng PKL & Guinot D (2008) Two new genera and
two new species of troglobitic false spider crabs (Crustacea:
We thank Rob Stuebing for making the arrangements to Decapoda: Brachyura: Hymenosomatidae) from Indonesia,
make collections at this site in Grand Perfect; and to Tan with notes on Cancrocaeca Ng, 1991. Zootaxa, 1739: 21–40.
Ng PKL (1988) The freshwater crabs of Peninsular Malaysia and
Heok Hui for collecting the specimens used for the study.
Singapore. Department of Zoology, National University of
The late Michael Türkay (SMF) kindly hosted the second Singapore, Shinglee Press, Singapore, viii + 156 pp., 63 figs.,
author in his visits to examine the type of A. kadamaiana. 4 colour pls.
We are also grateful to Keith Christenson and Tan Heok Hui Ng PKL (1989) The identity of the cavernicolous freshwater crab
for allowing us to use their photos of A. rhadamanthysi and Potamon (Thelphusa) bidiense Lanchester, 1900 (Crustacea:
A. sarang, respectively. We thank Jose Christopher Mendoza Decapoda: Brachyura: Gecarcinucidae) from Sarawak, Borneo,
and Célio Magalhães for their constructive comments of with description of a new genus. Raffles Bulletin of Zoology,
this manuscript. The first author thanks the Institute of 37(1–2): 63–72.
Biodiversity and Environmental Conservation, Universiti Ng PKL (1991) Bornean freshwater crabs of the genus
Arachnothelphusa gen. nov. (Crustacea: Decapoda: Brachyura:
Malaysia Sarawak, for laboratory space and technical support.
Gecarcinucidae). Zoologische Mededelingen, 65: 1–12.
Research Permit No. NCCD.907.4.4 (JLD. 13)-307 was Ng PKL (2013) Stygothelphusa cranbrooki, a new species of cave
granted by the Sarawak Forest Department. The research crab from Gua Sireh, Sarawak, Malaysia (Crustacea: Decapoda:
is partly supported by Project No. GL(I01)/MPOB/03/2016. Brachyura: Gecarcinucidae). Raffles Bulletin of Zoology,
Supplement 29: 91–97.
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